J. Hydrol. Hydromech., 65, 2017, 4, 333–342 DOI: 10.1515/johh-2017-0048
Drought impact on ground beetle assemblages (Coleoptera, Carabidae) in Norway spruce forests with different management after windstorm damage – a case study from Tatra Mts. (Slovakia)
Zbyšek Šustek1*, Jaroslav Vido2, Jana Škvareninová3, Jaroslav Škvarenina2, Peter Šurda4
1 Institute of Zoology of Slovak Academy of Sciences, Dúbravská cesta 9, SK–845 06 Bratislava, Slovakia. 2 Department of Natural Environment, Faculty of Forestry, Technical University in Zvolen, T. G. Masaryka 24, SK–960 53 Zvolen, Slovakia. E-mail: [email protected] 3 Department of Applied Ecology, Faculty of Ecology and Environmental Sciences, Technical University in Zvolen, T.G. Masaryka 24, SK–960 53 Zvolen, Slovakia. 4 Institute of Hydrology SAS, Dúbravská cesta 9, 841 04 Bratislava, Slovakia. E-mail: [email protected] * Corresponding author. Tel.: +421 2 59 30 26 17. E-mail: [email protected]
Abstract: After the windstorm of November 2004, the ground beetle assemblages (Coleoptera, Carabidae) differentiat- ed after the windstorm into four groups reflecting degree of damaging and forestry management (intact stand, fallen timber in situ, extracted timber, fire). The stand with fallen timber reduced abundances of original species. Removal of timber eliminated sensitive forest species and favored tolerant species, whereas the fire allowed invasions of field spe- cies. Later, the assemblages on burned sites converged to those in the unburned sites. Their restoration has a sigmoid-like course. Independently on the above differentiation and course assemblage succession, episodes of severe drought resulted with a 1–2-years delay in sudden decline of number of individuals and species. Their numbers were restoring after longer humid periods. Because these extremes occur with a considerable regularity, the observed extremes of fluctuations of number of species and individuals represent the variability limits of the Carabid assemblages in such conditions. The Standardized Precipitation Evapotranspiration Index was shown, using the cross-correlation of SPEI and number of indi- viduals and species of Carabids, as a suitable means to explain and predict such changes for the period of 1–2 years.
Keywords: Drought; Ground beetle; Windstorm; Forest management; Norway spruce.
INTRODUCTION artificial restoration of the ecosystem (Šustek, 2009, 2013; Šustek and Čejka, 2009; Šustek and Vido, 2013). The second The Tatra Mountains – the highest mountain range of the type of changes was common as for the assemblages in intact West Carpathians have a specific climatic, orographic and stands as for those in differently damaged and restoring stands. ecological character. Because of this, unique fauna and flora They included simultaneous declines or increases in number of have been formed there. However, because of the mountain individuals, species and biomass in one-year samples. The orography, damaging downslope wind called as Bora occurs eight-year investigation indicated that they have periodical periodically (Fleischer et al., 2009). This winds cause severe character correlated with the occurrence of short termed damages on forest ecosystems repeatedly on the same places drought episodes (Šustek and Vido, 2013). In addition there and influence structure of the biocoenoses. For the first time was observed a slight trend of spreading species with occur- they were described and mapped by Mrkos (in Gregor, 1929). rence optimum in lower altitudes and increase of representation The last Bora downslope wind occurred in November 2004. of species having the lower limit of altitudinal distribution in This was a huge devastating catastrophe that damaged almost lowlands (Lővei, 2008). 25% of the Norway spruce forests on southern slopes of High The aims of this paper are: (1) to describe the periodic and Tatra. In addition, in 2005 a considerable part of this area was non-periodic changes in mountain Carabid assemblages linked affected by a wide-spread forest fire. It had a profound impact to different forestry management and, first of all, occurrence of on the biota (Mezei et al., 2014a, 2014b; Renčo et al., 2015; short-term drought episodes and; (2) to compare the suitability Šustek and Vido, 2013; Urbanovičová et al., 2013). The ecosys- of the Standardized Precipitation Evapotranspiration Index for tems disturbed in this way are more prone to hydro- 6 and 12 months, using the cross-correlation, to explain or even meteorological hazards, in particular to drought (Ježík et al., predict changes in Carabid assemblages for about 1–2 years. 2015; Kurjak et al., 2012). Because of the wide concept of ecosystem, it is necessary to study impacts of drought using MATERIAL AND METHODS proper bioindicators. The Carabids (ground beetles) are espe- Study area cially suitable for this purpose because of their enormous eco- logical differentiation, high sensitivity to humidity changes and The investigations were carried out in six study plots repre- limited mobility (Lővei, 2008). senting an intact Norway spruce stand and a habitat wit fallen The Carabid assemblages in mountain forests ecosystems of timber in situ, two habitats with extracted timber and two habi- High Tatra damaged in November 2004 by the wind catastro- tats with extracted timber additionally damaged by large scale phe showed two types of changes. The first type of changes was fire in July and August 2005. The plots were selected by the staff connected with destruction of the stands, the subsequent human of the Investigation Station of the High Tatra National Park activities in the damaged area and with the spontaneous or (Fleischer, 2008) to coordinate the international investigations
333 Zbyšek Šustek, Jaroslav Vido, Jana Škvareninová, Jaroslav Škvarenina, Peter Šurda
Table 1. Survey of study plots in the area affected by the windstorm in High Tatra on 19 November 2004.
Locality Vyšné Hágy Tatranská Tatranská Nový Tatranské Tatranské reference plot Lomnica, Jamy, Polianka, Smokovec, Zruby Zruby Danielov dom Vodný les lower plot upper plot Locality REF NEXT EXTd EXTl FIRl FIRh abbreviations Geographical 49°07′17.5″N 49°09'33.7"N, 49°07′15.3″N 49°08'07.6"N, 49°07′49.3″N 49°08′02.7″N coordinates 20°06′15.0″E 20°15'07.9" E 20°09′46.0″E 20°12'24.8" E 20°11′49.1″E 20°11′30.1″E Altitude [m] 1233 1062 1060 1022 1015 1095 Vegetation zone Spruce Spruce Spruce Spruce Spruce Spruce Trophical series Acidophilou Acidophilou Acidophilou Acidophilou Acidophilou Acidophilou mesophilous mesophilous mesophilous mesophilous mesophilous mesophilous Forest type Sorbi Sorbi Sorbi Sorbi Sorbi Sorbi Piceeta Piceeta Piceeta Piceeta Piceeta Piceeta Degree of Intact Timber Timber Timber Timber Timber damaging mature in situ extracted, extracted, extracted, extracted, spruce forest unburned unburned burned burned started after the windstorm. In this stuffy, the sites (Table 1) are Methods described according to the Zlatník´s phytocoenological system of forests ecosystems (Raušer and Zlatník, 1966; Zlatník, Occurrence of drought episodes are characterized by The 1976). Standardized Precipitation Evapotranspiration Index (SPEI) Mountain climate of spruce vegetation zone is characterized (Hayes et al., 1999; Vincente-Serrano, 2010) calculated for the by a short growing season (from 70 to 100 days) and excess period 1960–2014 based on the data from the meteorological precipitation (Hlavatá et al., 2011). According to the measure- station Tatranská Lomnica. SPEI is logically based on calcula- ments at the stations of the Slovak Hydro–Meteorological Insti- tion principle of the Standardized Precipitation Index (McKee tute (SHMI), mean annual precipitation total fluctuates between et al., 1993). However the main advantage comparing to the 870 and 965 mm, and from 550 to 575 mm in the growing SPI is that the SPEI calculates balance between precipitation season (April to September). Mean annual totals of potential and potential evapotranspiration. In 99%, the SPEI values move evapotranspiration are in the range from 420 to 445 mm. Mean within the limits –3 and +3 and basing on the cumulative prob- annual air temperature is from 3.6 to 4.6°C, and from 9.0 to ability distribution the concrete values can be interpreted by 10.4°C in the growing season. January is the coldest month means of the Table 2. Thus this interpretation indicates signifi- (–5.3°C), and July is the warmest month (13.8°C). The ampli- cance of the drought episode that means a period with continu- tude of air temperature (19.1°C) indicates the interior, montane ous occurrence of negative values of SPEI. continentality. Snow cover lasts for about 110 to 155 days, and its average height is between 88 and 180 cm. Climatically, the Table 2. Cumulative probability distribution of the SPEI. studied region is classified as a humid cool climatic region, and a cool mountainous subregion (Hlavatá et al., 2011; Lapin et Values Character Number of occurrence al., 2002). of SPEI of deviation of situations within 100 years Natural montane Norway spruce (Picea abies) vegetation ≥ 2.0 Extremely humid 2.5 zone is a typical example of the extrazonal occurrence of the 1.5 to 1.99 Very humid 5 boreal taiga biome in the nemoral zone of the European moun- 1.0 to 1.49 Medium humid 10 tains (Plesník, 2004; Škvarenina et al., 2004). A forest type of –0.99 to 0.99 Close to normal 66 –1.0 to –1.49 Medium dry 10 Sorbi Piceeta represents communities where spruce is a domi- –1.5 to –1.99 Very dry 5 nant tree species. Admixed tree species are European larch ≤ –2.0 Extremely dry 2.5 (Larix decidua), and Swiss stone pine (Pinus cembra). Both larch and Swiss pine are more abundant on rocky soils, where For purposes of our study we calculated continuous SPEI for spruce cannot compete with them. Larch as a relatively short- 12 months (further SPEI-12). The SPEI-12 has been also used living (successional) tree species occurs also on deeper, loamy to uncover the influence of winter precipitation regime on soils, in particular at sites with frequent wind-throws. From spring ecological response of the spruce ecosystem in the High coniferous tree species, the Scotch pine (Pinus sylvestris) oc- Tatra Mountains. SPEI-12 has been cross-correlated (using the curs individually in the forests. Broadleaved tree species natu- PAST program, Hammer, 2012) with number of individuals rally occur only sparsely, in particular at sites with open stand and of species of ground beetle assemblages on the localities canopy of spruce forests (after the disturbance by wind, insects, described above. but also due to ageing). From broadleaved tree species, rowan The beetles were pitfall trapped. Six formalin traps (0.75 l (Sorbus aucuparia), and Carpathian hairy birch (Betula pu- plastic jars with 90 mm opening) were exposed in a line in 10 bescens ssp. carpatica) are frequent, and at moister, nutrient- m distances in each plot from end of May until early November richer sites we can also find the sycamore maple (Acer pseudo- 2007–2014 and emptied approximately in on month intervals. platanus). In this part of the Tatras, the spruce vegetation zone The beetles from each set of traps were summed to obtain one- forms the upper timberline. Above the continuous spruce stands season samples (see Supplementary material) that were used at there is a zone of open stands, tree groups and individual trees, further analyses. Scientific names of species are adopted ac- often mixed with dwarf mountain pine (Pinus mugo) cording to Hůrka (1996). The habitat and humidity preference (Škvarenina et al., 2004; Zlatník, 1976). of each species (Table 3) was characterized by semi- quantitative scales elaborated by Šustek (2004) basing on litera- ture (Burmeister, 1939; Desender, 1986a, b, c, d; Freude et al.,
334 Drought impact on ground beetle assemblages
1976; Lindroth, 1949; Lővei, 2008; Sharova, 1981; Šustek, Table. 3. Scientific names of species and characteristics of their 1992, 1994a, 1994b; Šustek, 2000; Thiele, 1977) and on the demands to vegetation cover (scale 1 – 4 discontinuous herbage author’s field experience in many ecosystems types in Central stratum, without wooden plants to complete shadowing by trees) Europe. The humidity scale is represented by eight degrees and humidity (scale 1 – 8 = strongly xerophilous to strongly hygro- ranging from 1 to 8 (1 = extremely xerophilous species of philous). The species are aggregated in major ecologic groups. steppe-like habitats, 4 = mesohygrophilous, 8 = extremely Vegetation Humidi- hygrophilous species of riverbank or swampy habitats), while Species cover ty the habitat preference by four degrees (1 = heliophilous species of open habitats, with discontinuous cover, 4 = stenotopic forest 1. Stenotopic forests species species preferring shadowing by completely closed canopy). Pterostichus nigrita (Fabricius, 1792) 4 8 These values were used to calculate the humidity preference Pterostichus strenuus (Panzer, 1797) 4 7 and vegetation cover preference indices of Carabid assemblage. Pterostichus niger (Schaller, 1783) 4 6 They were calculated as the average preference of all species in Carabus coriaceus (Linnaeus 1758) 4 5 one-year samples weighted by number of individuals of each Carabus glabratus (Paykull, 1790) 4 5 species as it is used in methods of direct ordination (Poole, Carabus linnei (Dejean, 1826) 4 5 1974). The structural changes of assemblages are quantified by Carabus violaceus (Linnaeus, 1758) 4 5 number of species and cumulative number of individuals in Cychrus caraboides (Linnaeus, 1758) 4 5 one-year catches (see Supplementary material). The ordination Leistus piceus (Frölich, 1799) 4 5 of the assemblages was carried out by non-parametric multidi- Leistus terminatus (Hellwig in Panzer, 1793) 4 5 mensional scaling (NMS) using the program PAST (Hammer, Pterostichus angustatus (Duftschmidt, 1812) 4 5 2012) and Horn´s index as a measure of proportional similarity. Pterostichus burmeisteri (Heer, 1801) 4 5 Pterostichus aethiops (Panzer, 1797) 4 5 RESULTS AND DISCUSSION Pterostichus foveolatus (Duftschmidt, 1812) 4 5 Pterostichus oblongopunctatus (Fabricius, 1787) 4 5 In the whole investigation period altogether 50 species were Pterostichus unctulatus (Duftschmidt, 1812) 4 5 recorded in all study plots (Table 3 and Supplementary materi- Trechus latus (Puzeys, 1847) 4 5 al). They include two sharply differing ecological groups – Trechus striatulus (Putzeys, 1847) 4 5 stenotopic forest species unable to fly and requiring permanent Trichotichnus laevicollis (Duftschmidt, 1812) 4 5 shadowing by closed tree canopy and heliophilous mostly well Carabus auronitens (Fabricius, 1792) 4 4 flying species bound to non-forests, natural and artificial eco- Carabus hortensis (Linnaeus, 1758) 4 4 systems and several species showing obvious preference for Carabus nemoralis (O. F. Müller, 1764) 4 4 forest, but tolerating substitution of closed forest vegetation Loricera caerulescens (Linnaeus, 1758) 4 4 with the high grassy stands, like Carabus violaceus and Cara- Molops piceus (Panzer, 1793) 4 4 bus glabratus (Magura, 2002). Among them, 7–21 species were recorded in individual plots and years. Their number was mod- 2a. Eurytopic species preferring open landscape erately positively correlated (r = 0.3148) with number of Calathus metalicus (Dejean, 1828) 3 5 trapped individuals that fluctuated from 22 to 376. The higher Calathus micropterus (Duftschmidt, 1812) 3 3 cumulative numbers of individuals have been obtained espe- Carabus arvensis (Herbst, 1784) 2 5 cially from abundant occurrence of small (2.4–7.0 mm, genera Agonum micans (Nicolai, 1822) 2 7 Microlestes, Bembidion) and medium sized species (7.0–18.0 Anisodactylus binotatus (Fabricius, 1792) 2 6 mm, especially genera Amara, Poecilus, Harpalus) (for body Agonum sexpunctatum (Linnaeus, 1758) 2 5 sizes of individual species see Freude et al. (1976) or Hůrka Europhilus gracilipes (Duftschmidt, 1812) 2 5 (1996)) predominating in the damaged plots with extracted Notiophilus biguttatus (Fabricius, 1779) 2 4 timber. In contrast of these patterns big species (16–40 mm, Notiophilus palustris (Duftschmidt, 1812) 2 4 almost exclusively genus Carabus) represented a major part of 2b. Strictly open landscape species the assemblages in the intact plot and in the plot with timber in Trechus amplicollis (Fairmair, 1859) 2 5 situ. Harpalus quadripunctatus (Dejean, 1829) 2 4 The between-year changes of number of species and cumula- Poecilus cupreus (Linnaeus, 1758) 2 4 tive number of individuals (Figs. 1–2) show a similar trend inde- Poecilus versicolor (Sturm, 1824) 2 4 pendently on the momentary degree of damaging the assemblage. Harpalus distinguendus (Duftschmidt, 1812) 1 4 In 2008, they suddenly dropped deeply under the level of the Harpalus latus (Linnaeus, 1758) 1 4 precedent year, but since 2009 they gradually increased. Num- Pseudoophonus rufipes (De Geer, 1774) 1 4 ber of species and individuals culminated in 2010 or 2011. The Harpalus affinis (Schrank, 1784) 1 3 extremely high values in 2011 in both burned plots are due to Amara aenea (De Geer, 1774) 1 3 invasion of the well flying Amara nitida (see Supplementary material). In 2012 numbers of individuals and species deeply Amara erratica (Duftschmidt, 1812) 1 3 dropped again and approximated their minimum values of 2008. Amara eurynota (Panzer, 1797) 1 3 In the next two years the numbers of species and individuals Amara familiaris (Duftschmidt, 1812) 1 3 stabilized at the approximately same level. However, in indi- Amara lunicollis (Schiodte, 1837) 1 3 vidual plots they show moderately different directions. In the Amara nitida (Sturm, 1825) 1 3 intact reference plot number of individuals continued to de- Amara ovata (Fabricius, 1792) 1 3 crease, but in one of the burned plots FIRE-l it increased. Num- Bembidion lampros (Herbst, 1784) 1 3 ber of species also moderately decreased on most plots, but it Microlestes maurus (Sturm, 1827) 1 2
slightly increased in the burned plot FIRE-h (see Supplementary
material).
335 Zbyšek Šustek, Jaroslav Vido, Jana Škvareninová, Jaroslav Škvarenina, Peter Šurda
25 Lag REF NEXT -4 -3 -2 -1 0 1 2 3 4 EXTd EXTv 1 20 FIRl FIRu 0.8 REF NEXT EXTd EXTv 15 0.6 FIRl FIRh
0.4
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0 -0.4 2007 2008 2009 2010 2011 2012 2013 2014 -0.6 400 -0.8 REF NEXT 350 EXTd EXTv -1 FIRl FIRh 300 Lag -4 -3 -2 -1 0 1 2 3 4 250 1 200 0.8 REF NEXT
Individuals 150 EXTd EXTv 0.6 FIRl FIRh 100 0.4 50 0.2 0 2007 2008 2009 2010 2011 2012 2013 2014 0
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Figs. 4–5. Cross-correlations of fluctuations of number of Carabid species (above) and individuals (bellow) with SPEI-12 months.
Figs. 1–3. Changes in number of species (1), cumulative number of individuals (2) and changes SPEI-12 (3) in six sites in High Tatra in 2007–2014 (abbreviations as in Table 1).
The changes described above coincided with course of changes of SPEI-12 (Figs. 1–3). In late 2006 and in 2007, there occurred a drought indicated by a sudden drop of this index. SPEI-12 was low in whole 2007 and rarely also 2008. In 2010, when number of species and individuals started to increase, SPEI-12 also showed high values ranging from 1.0 and 2.0. Occurrence of high values of this index is the more continuous; Fig. 6. Long-termed fluctuations of SPEI 12 months in High Tatra the longer periods are represented by it (Fig. 3). SPEI-12 in in 1960–2013. 2012 declined to the interval 0.0 to –1.0. This drop anticipated the strong decline of number of species and individuals in individuals. After a prolonged increase of SPEI-12 to the level 2012–2014 (Figs. 1–2). 1.0 to 2.0, a strong increase of number of individuals and spe- In 2012 and 2013, SPEI-12 declined to –2.0. In 2014 this in- cies followed. dex started to increase again. This moderate increase coincides The fluctuations in number of species and individuals are with increase of number of individuals of Carabids in some cross-correlated with fluctuations in SPEI-12 months. The plots in 2014 (Figs. 1–2). Comparison of course of changes in maximums of the cross-correlation coefficients occur mostly number of individuals and species with occurrence of drought with a 0–2-year lag (Figs. 4–5). periods indicated by SPEI index show that changes in both For prognosis of development of the cumulative number of number of species and individuals mostly occur with an ap- individuals of Carabid assemblages basing on climatic proximate delay of 1–2 years after incidence of extreme fluctuations it seems that the SPEI-12 has a considerable drought or rainy years. indicative value. The similar, but slightly moderately phase Figures 1–2 show that a sudden decline of number of species shifted course of fluctuation of the climatic factors and of and individuals of Carabids follows after a longer occurrence number of individuals and species in 2007–2012 puts question of values of SPEI close to –1.0 or lower. On contrary, a longer of periodicity of climatic fluctuation and their influence on occurrence of values around 0 already anticipate a moderate animal communities. Changes of SPEI-12 (Fig. 6) calculated increasing of number of species and cumulative number of for meteorological station Tatranská Lomnica for 1961–2014
336 Drought impact on ground beetle assemblages
show that fluctuations of SPEI-12 in range of –1.5 to +1.5, or 4.5 rarely even in the range of –2.0 to +2.0 occur regularly within 4 3–6 years, similarly as they did in the studied period 2007– 2014. The long-termed fluctuations of SPEI and short-termed 3.5 reactions of Carabids allow to make a hypothesis concerning 3 long-termed fluctuations of Carabid populations and 2.5 assemblages. 2 At the same time the observed maximum and minimal val- 1.5 REF NEXT ues of number of individuals and species probably represent EXTd EXTv limits of variability of the Carabid assemblages in similar 1
Vegetation cover preference index FIRl IRFh mountain ecosystems (Figs. 1–2). However, from the viewpoint 0.5 of zoocoenological classification the Carabid assemblages, the 0 observed values can be indicative just for the assemblages in 2007 2008 2009 2010 2011 2012 2013 2014 forests on oligotrophic acid crystalline substrates, while in the 5 ecosystems on the polytrophic nitrogenous or basic substrates, the extremes will be situated at a higher level (Šustek, 2009) and their establishing needs an extensive field research in future. 4.5 The phase shift (lag) of fluctuations in Carabid assemblages after the climatic fluctuations has two different, but closely connected reasons. The shortage of humidity reduces activity of 4 all edaphic organisms. Thus it simultaneously reduces the mo- mentary activity of adult Carabids and their chance to mate and REF NEXT Humidity preferenceHumidity index 3.5 EXTd Extv lay eggs, as well as chance of adults and larvae to find enough FIRl FIRh prey, to complete the development and to survive. Therefore the effect of drought is combined and occurs at several levels. 3 2007 2008 2009 2010 2011 2012 2013 2014 The Carabids are monovoltine, with two principal reproduction types in the holarctic region – the spring breeders mating and laying eggs in spring, where the new generation hibernates as Figs. 7–8. Changes in vegetation cover preference index (above) pupae or adults, and the summer breeders mating and laying and humidity preference index (bellow) of Carabid assemblages in eggs in late summer or early autumn, where new generation six sites in High Tatra in 2007–2014. hibernates as larvae (den Boer and den Boer-Daanje, 1990; Lővei, 2008; Thiele, 1977). There also exist a plastic reproduc- in situ (NEXT Tatranská Lomnica Jamy). In all three cases the tion type, but it is represented just by very few species, like growth of their representation has a sigmoid character and the Pterostichus melanarius (de Boer and de Boer-Daanje, 1990). values asymptotically approximate to the level in the intact plot, In lowlands, with long growing season, species of both repro- irrespectively of two periods of extreme drought, which could duction types are represented in assemblages in an approxi- only temporarily inhibit restoration of the damaged ecosystems mately balanced proportion. Different timing of their reproduc- by reducing number of individuals and species. tion reduces competition pressures of species and forms clear At the same time, succession of Carabid assemblages in all seasonal aspects of Carabid assemblages in some ecosystems. damaged plots had a convergent character and reducing differ- In mountain conditions, the spring breeders predominate to ences between assemblages from the burned plots and other effectively use the short growing season that can be even insuf- plots with extracted timber. Before all, representation of the ficient for the complete development of one generation. Thus open landscape species (Poecilus cupreus, Poecilus versicolor, development of some species can be prolonged on two growing Amara spp.) declined in the burned plots (Fig. 9–10) while the seasons and generation can overlap. Under such circumstances, more tolerant forest species (Carabus violaceus, Carabus gla- the extreme drought in a growing season or even in a short bratus, Molops picues) gradually spread here (Figs. 10–13, see period can essentially inhibit development of next generation, also Supplementary material). It was allowed by emergence of but with an impact visible as late as in following growing sea- pioneer wooden vegetation (Populus tremula, Sambucus race- son, if the beetles are monitored using pitfall traps. On contrary, mosa, Salix caprea, Betula alba) providing at least local shad- the restoration of the decimated population will need a longer owing, as well as by change of herbage stratum due to mowing period of normal or increased humidity, as it was observed in the extensive stands of Chamerion angustifolium and it’s re- the years 2009 and 2010. placing by the grassy stands (especially Calamgrostis epigeios) The extreme drought may inhibit, while the increased hu- The overall course of this process is illustrated by ordination of midity may speed up restoration process of assemblages in the one-year samples from all studied plots (Fig. 14). The first axis damaged area. As a result the assemblages will slowly return to represents gradient of damaging decreasing from left to right. their original state. However, the climatic fluctuations little The second axis shows progress of succession of assemblages influence the momentary functional structure of Carabid as- on damaged plots from 2007 to 2014. In early stages the semblages and direction of their succession (restoration). It is assemblages on the burned plots differed from unburned plots clearly shown by relative representation of species requiring the with extracted timber by pulse-like invasions of species charac- permanent shadowing (Fig. 7) and higher humidity (Fig. 8). In teristic for arable land, especially by Poecilus cupres (later spite of incidence of the periodic, climatically conditioned replaced by Poecilus versicolor), Pseudophonus rufipes changes in number of species and of individuals (Figs. 1–2), and several species of genus Amara (see Supplementary mate- representation of these two major groups of species (Table 3) rial). There also occurred heliophilous species Microlestes minu- increased in all damaged plots with extracted timber and tus and Bembidion lampros that prefer sites with discontinuous, converged to their stable representation in the intact reference plot patchy-like herbage vegetation arising during cutting of herbage (REF Vyšné Hágy) and in the damaged plot with fallen timber vegetation. At the end of the investigation period, all assemblages
337 Zbyšek Šustek, Jaroslav Vido, Jana Škvareninová, Jaroslav Škvarenina, Peter Šurda
80 REF NEXT 70 EXTd EXT v 60 FIR l FIR h
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10 Fig. 14. NMS ordination (Horn´s index) of one-years samples of 0 Carabids in six sites in the windstorm area in High Tatra in 2007– 2007 2008 2009 2010 2011 2012 2013 2014 2014 (abbreviations H – reference plot in Vyšné Hágy [= REF], J – plot with fallen timber in situ in Jamy [= NEXT], D – unburned Figs. 9–10. Changes in relative abundance of two most abundant plot with extracted timber near Danielov dom [= EXTd], V – un- open landscape species in six differently damaged and managed burned plot with extracted timber Vodný les Zd and Zh – lower sites in High Tatra in 2007–2014 (symbols as in Table 1). and upper burned plot with extracted timber in Tatranské Zruby [= FIRl and FIRh]). The arrows show succession direction of as- semblages in the burned plots. The first axis shows damage degree, 80 REF NEXT the second axis shows direction of succession from 2007 towards 70 EXTd EXT v 2014. s 60 FIR l FIR h 50 from the plots with extracted timber form a common cluster 40 that shifts to right side of the ordination space, towards to the 30
% of C. glabratu % assemblages from the intact plot and the plot with timber in 20 situ. However, the complete restoration of assemblage in the 10 damaged plots will be finished in a relatively remote future. 0 2007 2008 2009 2010 2011 2012 2013 2014 First of all the stenotopic forest Carabus linnei, C. auronitens, Cychrus caraboides, Leistus piceus, Pterostichus unctulatus, P.
burmeisteri, P. foveolatus, Calathus micropterus are missing in 80 them or occur there only exceptionally. Just these species repre- 70 REF NEXT EXTd EXT v sent the specific component of the assemblages in intact refer-
s 60 FIR l FIR h ence plot. To certain degree they also survive in the plot with 50 timber in situ (see Supplementary material). 40 In the last years, there also appeared other trend in all as- 30 semblages – increasing portion of two species having optimum % of C. violaceu C. % of 20 of distribution in highlands or lower limit of (Carabus vio- 10 laceus, Carabus glabratus) and showing a considerable toler- 0 2007 2008 2009 2010 2011 2012 2013 2014 ance to deforestation also in other regions (Magura, 2002). In the plot EXTl Carabus coriaceus, C. hortensis and C. nemoralis having optimum of vertical distribution in lowlands 80 started to occur in 2009–2011 (Fig. 15 and Supplementary 70 REF NEXT EXTd EXT v material). Thus the assemblage structure slowly shifts toward 60
s FIR l FIR h the assemblages that are characteristic of lower altitudes 50 (Šustek, 2014). However, interpretation of spreading of these 40 three species is not simple, because it started and furthermore is 30 concentrated in the plot at western margin of the Nový Smo- % of M. piceu of M. % 20 kovec town, where a temperature island may occur and where 10 Carabus coriaceus was observed one year before its recording 0 in this plot. 2007 2008 2009 2010 2011 2012 2013 2014 The obtained data indicate that three processes run simulta- neously in Carabid assemblages in the windstorm area in High Figs. 11–13. Changes in relative abundance of three tolerant forest Tatra: (1) overall periodic changes connected with the short- species in six differently damaged and managed sites in High Tatra termed, more or less regularly occurring climatic fluctuations, in 2007–2014 (symbols as in Table 1).
338 Drought impact on ground beetle assemblages
ini, Panageini, Odacanthini, Masoreini, Lebiini, Brachinini), Brussel, 48 p. Fleischer, P., 2008. Windfall research and monitoring in the 25 High Tatra Mts., objectives, principles, methods, and current
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Dominance in % Dominance 5 M., Škvarenina, J., Holecy, J. (Eds.): Bioclimatology and Together 0 Natural Hazards. Springer, Netherlands, pp. 145–154. C. coriaceus 2007 Freude, H., Harde, W., Lohse, G.A., 1976. Die Käfer Mitteleu- 2008 C. nemoralis 2009 2010 ropas, Band 2, Adephaga 1 – Goecke and Evers, Krefeld, 2011 C. hortensis 2012 2013 302 p. 2014 Gregor, A., 1929. Povětrnost (Počasí a podnebí) našeho státu [Weather and climate of Czechoslovakia]. In: Dědina, V., Fig. 15. Spreading of three thermophilous species of the genus Slavík, F., (Eds.): Československá vlastivěda [Geography of Carabus with optimum of vertical distribution in lowlands in the Czechoslovakia]. Příroda „Sphinx“, Praha, pp. 176–236. (In plot EXT l. Czech.) Hammer, O., 2012. PAST: Paleontological Statistics, Version (2) non-periodic changes connected with spontaneous or artifi- 2, 16, Reference manual. Natural History Museum, Univer- cial restoration of plant cover and Carabid assemblages in the sity of Oslo, Olso. damaged plot and (3), at present relatively slight, but probably Hayes, M., Svoboda, M., Wilhite, D., Vanyarkho, O., 1999. also long-termed changes resulting from the moderate warming Monitoring the 1996 Drought using the Standardized Precip- of the climate and spreading of species with distribution opti- itation Index. Bull. Amer. Meteor. Soc., 80, 429–438. mum at lower altitudes. Hůrka, K., 1996. Střevlíkovití České a Slovenské republiky, Although the studied localities are situated in a 12 km long Carabid beetles of the Czech and Slovak Republic. Kabou- strip and difference of altitude of the lowest and highest plot is rek, Zlín, 565 p. (In Czech and English.) approximately 250 m, values SPEI-12 calculated on the base of Hlavatá, H., Škvarenina, J., Čepčeková, E., 2011. Bio- data from a single meteorological station (NEXT Tatranská klimatické a zrážkové pomery v horských a vysokohorských Lomnica Jamy), situated at the eastern part of the study area, oblastiach Slovenska na príklade Tatier [Bioclimatic and are sufficiently representative for characterizing climatic condi- precipitation conditions in mountain and alpine areas of Slo- tion for existence of Carabid assemblages in the whole studied vakia on example of High Tatras Mts.]. Životné prostredie, area. 45, 64–70. (In Slovak with English abstract.) Ježík, M., Blaženec, M., Letts, M.G., Ditmarová, Ľ., Sitková, Acknowledgements. The authors gratefully acknowledge the Z., Střelcová, K., 2015. Assessing seasonal drought response Grant agency VEGA for the financial support by the grants of Norway spruce (Picea abies (L.) Karst.) by monitoring 2/4068/04, 2/7079/27, 2/0140/10, 2/0101/14, 1/0589/15, stem circumference and sap flow. Ecohydrology, 8, 378– 1/0367/16, 1/0463/14, APVV-15-0425 and APVV-15-0497 and 386. project KEGA 017TU Z-4/2016. Kurjak, D., Střelcová, K., Ditmarová, Ľ., Priwitzer, T., Kmeť, J., Homolák, M., Pichler, V., 2012. Physiological response REFERENCES of irrigated and non-irrigated Norway spruce trees as a con- sequence of drought in field conditions. Eur. J. For. Res., Burmeister, F., 1939. Biologie, Ökologie und Verbreitung der 131, 1737–1746. europäischen Käfer auf systematischer Grundlage, I. Band: Lapin, M., Faško, P., Melo, M., Šťastný, P., Tomlain, J., 2002. Adephaga, I. Familiengruppe: Caraboidea. Hans Goecke Climatic regions, Landscape atlas of the Slovak Republic. Verlag, Krefeld, 207 p. Ministry of Environment of the Slovak Republic and Slovak den Boer, J.P., den Boer-Daanje, W., 1990. On life history Environmental Agency, Bratislava, Banská Bystrica, 95 p. tactics in Carabid beetles: are there only spring and autumn Lindroth, K., 1949. Die Fennoskandischen Carabidae, Eine breeders? In: Stork, N.E. (Ed.): The role of ground beetles in tiergeographische Studie. Wettetrgten and Kerbers Förlag, ecological and environmental studies. Intercept Ltd., Ando- Götteborg, 712 p. ver, pp. 247–256. Lővei, G., 2008. Ecology and conservation biology of ground Desender, K., 1986a. Distribution and ecology of Carabid bee- beetles (Coleoptera, Carabidae) in an age of increasing hu- tles in Belgium (Coleoptera, Carabidae) Part 1, Species 1–80 man dominance.
339 Zbyšek Šustek, Jaroslav Vido, Jana Škvareninová, Jaroslav Škvarenina, Peter Šurda
Mezei, P., Grodzki, W., Blaženec, M., Škvarenina J., Geobiocenologická typizace krajiny a její aplikace [Geobio- Brandýsová, V., Jakuš, R., 2014b. Host and site factors af- cenological landscape typology and applications]. Geobio- fecting tree mortality caused by the spruce bark beetle (Ips cenologické spisy, 9, 210–214. typographus) in mountainous conditions. Forest Ecol. Man- Šustek, Z., 2009. Changes of secondary productivity of carabid age., 331, 196–207. communities (Insecta: Coleoptera) in natural forest ecosys- Plesník, P., 2004. Všeobecná biogeografia [General biogeogra- tems in relation to geological substrate and vertical zonality. phy]. Univerzita Komenského, Bratislava, 428 p. (In Slovak.) Oltenia, Studii şi comunicări Ştiinţele Naturii Muzeul Olten- Poole, R.W., 1974. An Introduction to Quantitative Ecology. iei Craiova, 25, 83–90. McGraw-Hill, New York, 532 p. Šustek, Z., 2013. Differentiation and succession of carabid Raušer, J., Zlatník, A., 1966. Biogeografie I [Biogeography I]. communities in the forests damaged by the wind catastrophe In: Svoboda, J., Stehlík, B. (Eds.): Atlas Československé in High Tatra in 2004. In: Friedl. M. (Ed.): Geobiocenologie socialistické republiky [Atlas of the Czechoslovak a její aplikace v lesnictví a krajinářství [Geobiocenology and Socialistic Republic], Praha, p. 21. its application in forestry and landscape engineering]. Geo- Renčo, M., Čerevková, A., Homolová Z., Gömöryová, E., biocenologické spisy, 15, 185–201. 2015. Long-term effects on soil nematode community struc- Šustek, Z., 2014. Shifts in representation of altitude preference ture in spruce forests of removing or not removing fallen Groups of Carabid Beetles (Coleoptera, Carabidae) in the trees after a windstorm. Forest Ecol. Manage. 356, 243–252. zone of windstorm calamite in High Tatra in November Sharova, K.I., 1981. Zhiznyenye formy zhuzhelits [Life forms 2004. In: Čelková, A. (Ed.): 21st International Poster Day of ground beetles]. Nauka, Moskva, 360 p. (In Russian.) Transport of Water, Chemicals and Energy in the Soil-Plant- Škvarenina, J., Križová, E., Tomlain, J., 2004. Impact of the Atmosphere System. Institute of Hydrology, Bratislava, pp. climate change on the water balance of altitudinal vegetation 335–346. stages in Slovakia. Ekológia, 23, Suppl 2, 13–19. Šustek, Z., Čejka, T., 2009. Coincidence of response of mollusks Šustek, Z., 1992. Windbreaks and line communities as migrati- (Mollusca) and ground beetles (Coleoptera, Carabidae) on wind ons corridors for Carabids (Col. Carabidae) in the agricultu- disaster in High Tatra in 2004. In: Hrubá, V., Štykar, J. (Eds.): ral landscape of South Moravia. Ekológia (ČSFR), 11, 259– Geobiocenologie a její aplikace v krajině [Geobiocenology and 271. their applications in landscape]. Brno, pp. 188–196. Šustek, Z., 1994a. Classification of the Carabid assemblages in Šustek, Z., Vido, J. 2013. Vegetation state and extreme drought as the floodplain forests in Moravia and Slovakia. In: Desen- factors determining differentiation and succession of Carabidae der, K. et al. (Eds.): Carabid Beetles, Ecology and Evolution, communities in forests damaged by a windstorm in the High pp. 371–376. Tatra Mts. Biologia, 68, 119–121. Šustek, Z., 1994b. Windbreaks as migration corridors for cara- Thiele, H.U., 1977. Carabid Beetles in Their Environments. A bids in an agricultural landscape. In: Desender, K. et al. Study in Habitat Selection by Adaptations in Physiology and (Eds.): Carabid Beetles, Ecology and Evolution, pp. 377– Behavior. Springer, Berlin, Heidelberg, New York, 330 p. 382. Urbanovičová, V., Miklisová, D., Kováč, Ľ., 2013. The effect of Šustek, Z., 2000. Spoločenstvá bystruškovitych (Coleoptera, Cara- windthrow, wild fire, and management practices on epigeic bidae) a ich využitie ako doplnkovej charakteristiky geobioce- Collembola in windthrown forest stands of the High Tatra Mts nologickych jednotiek: problémy a stav poznania [Community (Slovakia). Biologia, 68, 941–949. Coleoptera, Carabidae and their use as an additive characteris- Vincente-Serrano, S., Lasanta, T., Garcia, C., 2010. Aridifica- tics geobiocoenological units: problems and state of tion determines changes in forest growth in Pinus halepensis knowledge]. In: Štykar, J., Čermák P. (Eds): Geobiocenologická forests under semiarid Mediterranean climate conditions. typizace krajiny a její aplikace [Geobiocenological landscape Agric. For. Meteorol., 150, 614–628. typology and applications]. Geobiocenologické spisy, 5, 18–30. Zlatník, A., 1976. Přehled skupin geobiocénů původně lesních a (In Slovak.) křovinných v ČSSR [Overview of groups of geobiocoenes, Šustek, Z., 2004. Characteristics of humidity requirements and originally forest and shrub formations in Czechoslovakia]. relations to vegetation cover of selected Centra-European Zprávy Geografického ústavu ČSAV, 13, 55–65. (In Czech.) Carabids (Col., Carabidae). In: Štykar, J. Čermák, P. (Eds.): Received 19 December 2016
Accepted 21 September 2017
SUPPLEMENTARY MATERIAL
Table S1. Survey of species and number of individuals caught in six study plots High Tatra in 2007–2014: reference plot and plot with timber in situ (years marked just by the last digit), considerable part of the data match to that published by Šustek and Vido (2013).
Species Vyšné Hágy - REF Jamy - NEXT 7 8 9 0 1 2 3 4 7 8 9 0 1 2 3 4 A. micans A. sexpunctatum A. aenea A. erratica 1 1 A. eurynota A. familiaris A. lunicollis A. nitida A. ovata A. binotatus B. lampros C. metalicus 1
340 Drought impact on ground beetle assemblages
C. micropterus 9 12 10 13 1 2 4 C. arvensis 1 C. auronitens 18 1 6 10 16 2 1 1 3 9 3 C. coriaceus C. glabratus 7 1 3 3 9 8 8 21 15 1 6 11 8 3 6 24 C. hortensis C. linnei 17 2 8 14 15 2 1 11 25 2 3 8 2 1 1 C. nemoralis C. violaceus 29 9 18 53 89 67 47 49 10 6 14 30 31 24 31 8 C. caraboides 8 2 3 4 4 1 3 3 33 E. gracilipes H. affinis H. distinguendus H. latus H. quadripunctatus 3 1 34 L. piceus 1 L. terminatus L. caerulescens M. maurus M. piceus 7 3 4 4 1 1 1 1 N. biguttatus 4 2 2 1 1 1 1 N. palustris 1 P. cupreus P. versicolor P. rufipes P. aethiops 31 1 3 9 411 91 4 11 512 P. angustatus P. burmeisteri 17 5 13 25 25 17 14 16 5 10 14 5 P. foveolatus 44 9 25 57 94 12 6 4 4 1 2 9 2 2 2 P. niger P. nigrita P. oblongopuncatus 1 11 1 P. strenuus P. unctulatus 208 35 159 186 47 27 13 8 25 8 16 29 8 7 1 T. amplicollis T. latus T. striatulus 8 2 1 T. laevicollis 1 2 3 1 1 3 2 1 6 4 Number of individ. 372 77 251 376 328 143 93 96 106 24 56 125 80 51 54 46 Number of species 13 10 13 15 17 10 10 7 9 7 10 15 13 9 9 7
Table S2. Survey of species and number of individuals caught in six study plots High Tatra in 2007–2014: unburned plots with extracted timber (years marked just by the last digit).
Species Danielov dom EXTd Vodný les EXTv 7 8 9 0 1 2 3 4 7 8 9 0 1 2 3 4 A. micans A. sexpunctatum 1 1 A. aenea 22 5 2 1 A. erratica 102 12 26 18 7 5 14 9 12 8 2 A. eurynota 1 6 2 2 1 1 A. familiaris 3 1 1 1 1 A. lunicollis 1 A. nitida 2322 1 1 2 A. ovata A. binotatus 11 B. lampros 11 C. metalicus C. micropterus C. arvensis C. auronitens 11 1 2 1 C. coriaceus 12 4 5 62 C. glabratus 47 1 11 35 5 54 34 5 1 3 2 13 18 35 23 C. hortensis 1 3 3 2 C. linnei 4 1 C. nemoralis 5 5 32 C. violaceus 40 18 23 36 78 62 23 23 3 7 4 15 34 12 19 32 C. caraboides 2 2 1 1 1 E. gracilipes H. affinis 1 2 H. distinguendus 1 H. latus 1 2 H. quadripunctatus 3 1 1 1 L. piceus L. terminatus 1 L. caerulescens 1 1 5121 M. maurus 1 M. piceus 1 11311 2 11 6 42 N. biguttatus 1 5 2 N. palustris 1 P. cupreus 1 3 2 6 3 2 1 2 3
341 Zbyšek Šustek, Jaroslav Vido, Jana Škvareninová, Jaroslav Škvarenina, Peter Šurda
P. versicolor 7 4513 4 1 1 P. rufipes 12 1 1 1 P. aethiops 1 4 112112 1 3 5 87 P. angustatus 3 2 P. burmeisteri 2 1 2 1 2 5 2 4 3 3 P. foveolatus 1 1 2 4 P. niger 1 21 P. nigrita 2 31 P. oblongopunctatus 1 1 1 2 P. strenuus 1 1 P. unctulatus 28 1 9 15 3 1 2 4 T. amplicollis 4 2 T. latus 1 T. striatulus 13 T. laevicollis 1 1 2 1 1 1 1 1 Number of indiv. 236 46 73 137 175 84 88 63 57 22 44 52 95 58 79 78 Number of species 15 9 12 20 15 9 10 6 20 7 18 21 19 11 9 10
Table S3. Survey of species and number of individuals caught in six study plots High Tatra in 2007–2014: burned plots with extracted timber (years marked just by the last digit.
Species Tatranské Zruby lower FIRl Tatranské Zruby upper FIRh 7 8 9 0 1 2 3 4 7 8 9 0 1 2 3 4 A. micans 1 A. sexpunctatum 1 1 1 A. aenea 1 2 6 2 3 4 1 1 A. erratica 6 4 3 5 1 4 1 8 2 2 4 1 1 2 A. eurynota 21 1 6 1 2 3 1 2 A. familiaris 1 1 1 2 A. lunicollis 1 A. nitida 112 5 1 103 3 A. ovata 2 A. binotatus 2 1 B. lampros 26 1 4 1 4 9 3 C. metalicus C. micropterus 1 1 C. arvensis C. auronitens 1 1 2 3 1 1 3 1 1 C. coriaceus 1 C. glabratus 8 2 5 3 9 16 11 47 4 1 4 9 9 7 24 C. hortensis 1 C. linnei 7 C. nemoralis C. violaceus 2 6 10 21 79 33 19 82 1 3 5 17 38 31 33 29 C.caraboides 1 E. gracilipes 2 1 3 H. affinis H. distinguendus H. latus 1 H. quadripunctus 2 2112 1 1 L. piceus L. terminatus L. caerulescens 1 1 1 M. maurus 1 2 4 2 2 M. piceus 2 12 34 61 13 1 2 1 11 10 20 2 N. biguttatus 4 3 6 2 1 1 N. palustris 1 1 P. cupreus 17 21 25 9 4 1 1 5 8 13 8 3 P. versicolor 11719282 74 5 7 3 P. rufipes 1 2 1 2 1 P. aethiops 7 1 1 P. angustatus P. burmeisteri 1 1 1 1 1 P. foveolatus 1 1 2 1 1 P. niger 316 8 P. nigrita P. oblongopunctus 3 1 P. strenuus P. unctulatus 1 2 7 8 2 6 1 1 T. amplicollis T. latus T. striatulus T. laevicollis 21 1 2 1 Number of individ. 108 41 65 55 352 130 132 156 50 28 38 53 254 62 73 67 Number of species 20 9 9 13 15 13 11 8 17 7 14 15 15 9 9 11
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